Anales del Jardín Botánico de Madrid
Vol. 66(1): 65-84
enero-junio 2009
ISSN: 0211-1322
doi: 10.3989/ajbm.2209
Synopsis and lectotypification of Solanum (Solanaceae)
species endemic in the West Indies
by
Sandra Knapp
Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. [email protected]
Abstract
Resumen
Knapp, S. 2009. Synopsis and lectotypification of Solanum
(Solanaceae) species endemic in the West Indies. Anales Jard.
Bot. Madrid 66(1): 00-00.
Knapp, S. 2009. Resumen y lectotipificación de las especies
endémicas de Solanum (Solanaceae) en las Antillas. Anales Jard.
Bot. Madrid 66(1): 00-00 (en inglés).
The West Indies (Bahamas, Greater and Lesser Antilles) is an area
of high species richness and more importantly, species-level endemism in many groups. Solanum (Solanaceae) is diverse in the
region with 25 endemic species (S. bahamense is near-endemic,
bringing number to 26). Nineteen of these taxa are single-island
endemics, and some have not been re-collected since the early
20th century. A synopsis of the species endemic to the West Indies is presented, with full synonymy and distribution; a single
representative specimen is cited for each. All accepted names
and synonyms are lectotypified where necessary; many of these
taxa were described from collections in Berlin (B) that are no
longer extant. Lectotypes newly designated here are illustrated.
Las Antillas (Bahamas, Grandes Antillas y Pequeñas Antillas) es
un área con gran riqueza florística y, lo que es más importante,
con alto nivel de endemicidad en muchos grupos. Solanum
(Solanaceae) es un género muy diverso en la región, que tiene
25 especies endémicas (S. bahamense es casi endémica, con lo
que el número se elevaría a 26). Diecinueve de estos táxones son
endemismos de una sola isla, y algunos no se han vuelto a colectar desde principios del siglo XX. Se presenta ahora una sinopsis
de las especies endémicas de las Antillas, con todos los sinónimos y su distribución y con cita de un espécimen representativo
de cada especie. Todos los nombres aceptados y sinónimos se
han lectotipificado cuando ha sido necesario; muchos de estos
táxones fueron descritos a partir de colecciones que se conservaban en Berlín (B) pero que han desaparecido. Los lectótipos
designados aquí están ilustrados.
Keywords: typification, historic collections, America, exploration, West Indies, Caribbbean, diversity.
Palabras clave: tipificación, colecciones históricas, América, exploración, Antillas, Caribe, diversidad.
Introduction
an on-line taxonomic resource, Solanaceae Source. One
of the goals of the PBI Solanum project is to designate
lectotypes for all Solanum names, helping to stabilise
nomenclature and facilitate further taxonomic research.
The West Indies are usually defined as the islands
of the Caribbean comprising the Greater Antilles
(Cuba, Hispaniola, Jamaica, Puerto Rico), the Bahamas, and the Lesser Antilles (Windward and Leeward Islands), but excluding the islands of Trinidad
and Tobago, which are of continental origin, and the
Dutch Antilles. The geological complexity of the West
Indies has meant that the region is a natural laboratory for studies of evolution and diversification (Ricklefs
& Bermingham, 2007). The region as a whole is of
Solanum (Solanaceae) is one of the ten most speciesrich genera of angiosperms. With approximately 1,500
species distributed worldwide on all continents except
Antarctica, Solanum encompasses a wide range of
habitats, from deserts to tropical rain forests, and
habits, from annual herbs to canopy trees. The genus,
like the family Solanaceae, is most diverse in the Americas, and centres of species diversity in Solanum are
found in the Andes of South America, Australia and
East Africa. As part of the collaborative project “PBI
Solanum: a world-wide treatment” (see Knapp & al.,
2004; http://www.nhm.ac.uk/solanaceaesource), descriptions of all species of Solanum together with details of types and nomenclature are being provided via
66
S. Knapp
Table 1. Solanum species recorded in the West Indies (see Knapp, in press; Acevedo checklist web address; Solanaceae Source).
Single island endemics are in bold (Cuba, 6; Hispaniola, 7; Jamaica, 3; Puerto Rico, 1; Virgin Islands, 1; St. Vincent, 1).
Solanum species
S. acropterum Dunal
S. aculeatissimum Jacq.
S. aethiopicum L.
S. americanum Mill.
S. bahamense L.
S. betaceum Cav.
West Indian distribution
Jamaica
Bahamas, Greater and Lesser Antilles (exotic)
Cultivated in Lesser Antilles
Bahamas, Greater and Lesser Antilles
Bahamas, Greater and Lesser Antilles; all but Hispaniola
Cultivated
S. bicolor Willd.
S. boldoense A.DC.
S. campechiense L.
S. capsicioides All.
S. carolinense L.
S. chamaeacanthum Griseb.
S. conocarpum A. Rich.
S. crotonoides Lam.
S. dendroicum O.E. Schulz & Ekman
S. diphyllum L.
Lesser Antilles
Cuba
Greater Antilles
Greater Antilles
Greater Antilles
Cuba
Virgin Islands
Hispaniola
Hispaniola
Cultivated
S. donianum Walp.
S. elaeagnifolium Cav.
Bahamas, Greater Antilles
Bahamas, Greater and Lesser Antilles (exotic)
S. ensifolium Dunal
S. erianthum D. Don
Puerto Rico
Bahamas, Greater and Lesser Antilles
S. glaucescens Zucc.
S. gundlachii Urb.
S. havanense Jacq.
S. hazenii Britton
S. jamaicense Mill.
S. lanceifolium Jacq.
S. lycopersicum L.
S. macrocarpon L.
S. maestrense Urb. & Ekman
S. mammosum L.
Cuba (possibly cultivated)
Cuba
Greater Antilles
Lesser Antilles
Greater and Lesser Antilles
Greater and Lesser Antilles
Cultivated
Cultivated
Cuba
Bahamas, Greater and Lesser Antilles
S. melongena L.
S. microphyllum (Lam.) Dunal
S. moense Britton & Wilson
S. myriacanthum Dunal
S. nigrescens M. Martens & Galeotti
S. nudum Dunal
S. orthacanthum O.E. Schulz
S. pachyneurum O.E. Schulz
S. pachyneuroides Amshoff
S. polyacanthos Lam.
S. polygamum Vahl
S. punctulatum Dunal
S. pyrifolium Lam.
S. quitoense Lam.
S. rostratum Dunal
Cultivated
Bahamas, Greater Antilles
Cuba
Greater Antilles (exotic)
Greater and Lesser Antilles
Bahamas, Greater and Lesser Antilles
Hispaniola
Cuba
Cuba
Hispaniola
Greater Antilles
Jamaica
Hispaniola
Cultivated in Greater Antilles
Virgin Islands (exotic)
S. rugosum Dunal
S. schlechtendalianum Walp.
S. schulzianum Urb.
S. seaforthianum Andrews
Greater and Lesser Antilles
Greater Antilles
Hispaniola
Bahamas. Greater and Lesser Antilles
World distribution
Cultivated; originally from Africa
Worldwide
Florida
Cultivated worldwide; originally from
South America
Northern South America
Mexico, Central America
Central and South America
North America
Mexico and Central America; widely
cultivated in tropics
SE USA, Mexico
Widespread invasive; originally from the
Americas
Widespread invasive; originally from the
Americas
Mexico
Mexico, Central and South America
Mexico, Central and South America
Mexico, Central and South America
Worldwide; originally from South America
Cultivated; originally from Africa
Widespread in cultivation; originally from
South America
Cultivated worldwide; originally from Asia
Mexico, Central and South America
Mexico, Central and South America
Mexico, Central and South America
South America
North America, Mexico and Central America;
widespread invasive
Mexico, Central and South America
Mexico, Central and South America
Northern South America, widespread
in cultivation
Anales del Jardín Botánico de Madrid 66(1): 65-84, enero-junio 2009. ISSN: 0211-1322. doi: 10.3989/ajbm.2209
West Indian Solanum
67
Table 1. (Continuation).
Solanum species
S. selleanum Urb. & Ekman
S. sisymbrifolium Lam.
West Indian distribution
Hispaniola
Greater and Lesser Antilles
S. tampicense Dunal
S. tetramerum Dunal
S. torvum Sw.
S. triste Jacq.
S. troyanum Urb.
S. tuberosum L.
Greater Antilles
Greater Antilles
Bahamas, Greater and Lesser Antilles
Lesser Antilles
Jamaica
Cultivated
S. umbellatum Mill.
S. urens Dunal
S. wendlandii Hook.f.
Greater Antilles
St. Vincent
Cultivated
S. woodburyi Howard
S. wrightii Benth.
Puerto Rico
Cultivated
composite character, with the Greater Antilles, Bahamas and Lesser Antilles each having different geological origins and histories (Iturralde-Vinent &
MacPhee, 1999). The Bahamas have their origins in
the North American tectonic plate, and are older and
more stable than the rest of the Archipelago (Mullins
& Lynts, 1977). The Greater Antilles are primarily of
Central American crust origin (forming to the west of
present day Central America and moving to the east
on the Caribbean plate), while the Lesser Antilles are
derived from a volcanic arc at the eastern edge of the
Caribbean plate (Iturralde-Vinent & MacPhee, 1999;
Ricklefs & Bermingham, 2007). The Caribbean region as a whole has its origin as islands no earlier than
the middle Miocene (Iturralde-Vinent & MacPhee,
1999), and the Caribbean tectonic plate has many
blocks giving many of the islands a composite geological character (e.g., Hispaniola, Jamaica). IturraldeVinent and MacPhee (1999) posit land connections to
northern South America (the GAARlandia hypothesis) during the Eocene-Oligocene transition, while
others (Hedges, 1996; Glor & al., 2005; Ricklefs &
Bermingham, 2007) suggest dispersal accounts better
for taxon distribution patterns observed, particularly
for vertebrates.
The West Indies is an area of considerable species
richness in Solanum, but more interestingly, is home to
many unusual and poorly known endemic species. Of
the 62 species registered from the region (see Knapp,
in press), 9 are introduced in cultivation, 28 are found
elsewhere in the Americas and 25 are endemic. If we
include Solanum bahamense L. as a near endemic (outside the West Indies it occurs only in Florida; it is included here, bringing the number of endemics to 26),
then more than 1/3 of the species in the region are
World distribution
Widespread invasive; originally from
South America
Florida, Mexico and Central America
Worldwide in tropics
Northern South America
Worldwide; originally from Andean South
America
Mexico, Central and South America
Widely cultivated; originally from Central
America
found only there. Of these, 19 are single island endemics (see Table 1), making the West Indies critically
important for the conservation of Solanum diversity.
The vast majority of these taxa have not been included
in any phylogenetic analysis, and Whalen (1984), in his
conspectus of the spiny solanums considered most of
them to be of uncertain affinities. A tentative division
into morphological assemblages can be made, however, this should be regarded as preliminary at best, and
many of these taxa are isolated both morphologically
and probably phylogenetically (see Table 2). Bohs
(pers. comm.) had considerable difficulties aligning
the plastid DNA sequences of S. polygamum, the unusual dioecious species from the Virgin Islands (see
Knapp & al., 1998), so that it was not included in any
analyses (Levin & al., 2006; Weese & Bohs, 2007). The
West Indian endemic Solanum species are indeed morphologically unusual, and have combinations of characters not found elsewhere in the genus. It is apparent
from morphology that radiations of Solanum have occurred within the West Indies, and that species present
there will have their sister taxa in the region rather than
outside. Phylogenetic analysis of these taxa will certainly contribute to our understanding of hypotheses
of Caribbean biogeography, but as most of these taxa
are rare and many have not been collected recently, exploration and re-collection is a priority. To this end,
this synopsis is presented in order to highlight a
promising area for future phylogenetic research. Taxa
recorded as West Indian endemic Solanum but now regarded as synonyms of more widely distributed species
are presented in Table 3.
Many of the Solanum species from the West Indies
were described by O.E. Schulz, who worked in Berlin
during the early part of the 20th century. Schulz did not
Anales del Jardín Botánico de Madrid 66(1): 65-84, enero-junio 2009. ISSN: 0211-1322. doi: 10.3989/ajbm.2209
68
S. Knapp
Table 2. Preliminary groupings for West Indian endemic Solanum.
Species group
Geminata clade (S. havanense species group of Knapp, 2008b)
Dulcamaroid clade
Brevantherum clade
Leptostemonum clade (S. torvum species group, Torva clade of
Levin & al., 2006)
Leptostemonum clade (S. bahamense species group;
Bahamense clade of Levin & al., 2006)
Leptostemonum clade (putative S. tetramerum species group,
part of Robustum clade sensu Levin & al., 2006)
Leptostemonum clade (putative S. schulzianum species group)
Leptostemonum clade (S. crotonoides species group, allied to
Torva clade fide Levin & al., 2006)
Leptostemonum clade (incertae sedis, perhaps related to
Robustum clade of Levin & al., 2006)
Incertae sedis
Table 3. Species of Solanum recorded as endemic to the West
Indies now regarded as synonyms of widespread taxa (for those
taxa now recognised as members of the genus Lycianthes see
Knapp, in press and http://botany.si.edu/Antilles/WestIndies/
catalog.htm).
Solanum species
S. antillarum O.E. Schulz
S. daturifolium O.E. Schulz
S. fragile Griseb.
S. guanicense Urb.
S. indecorum A. Rich.
S. marmeladense Urb.
& Ekman
S. mucronatum Urb.
S. riedleii Dunal
S. sagreanum A. Rich.
Accepted name
S. nudum Dunal
S. torvum Sw.
S. schlechtendalianum Walp.
S. campechiense L.
S. americanum Mill.
S. campechiense L.
Cordia alliodora (Ruiz & Pav.)
Oken (Boraginaceae)
Solanum schlechtendalianum
Walp.
S. glaucescens Zucc.
cite herbaria for any of the collections he cited, so it is
generally assumed that he examined sets of material of
these plants at B. The specimens he examined in B are
no longer extant, necessitating the lectotypification of
all of his names, including those he coined in collaboration with Eric Ekman. Fortunately duplicates of
many of these collections are widely distributed, and
for the names associated with Ekman collections, the
top set of this material, annotated by Ekman, is kept at
S. I have been unable to trace duplicates of some of
the collections used by Schulz, but none of these
species are accepted names. In these cases I have not
neotypified these names here in the hope that future
searches will uncover duplicates of the collections
upon which these epithets are based. I have not lectotypified Lamarck’s names for Caribbean solanums, as
Putative component taxa
S. acropterum, S. conocarpum, S. havanense, S. troyanum
S. boldoense, S. pyrifolium
S. punctulatum
S. orthacanthum
S. bahamense, S. ensifolium, S. polyacanthos
S. microphyllum, S. tetramerum
S. chamaeacanthum, S. gundlachii, S. pachyneuroides,
S. schulzianum, S. selleanum, S. urens
S. crotonoides, S. dendroicum, S. moense, S. pachyneurum,
S. woodburyi
S. polygamum
S. maestrense
these are represented in the Lamarck herbarium in P
with single, clearly annotated sheets that link unambiguously with the protologues. Other names for
which the authors unambiguously cited herbaria
where they had seen material (e.g., Dunal often cited
G-DC or G-Boiss. [now integrated into G] explicitly
in his 1852 Prodromus treatment) have also not been
lectotypified.
Reasoning for the choice of all lectotypes is given,
along with the distribution of each species and a representative specimen from each island where it occurs. Where lectotypes have been chosen previously
full reference is given, but reasoning for the choice not
repeated here. Descriptions of all these species, discussions of their putative relationships, full specimen
citations and distribution maps can be found on Solanaceae Source (http://www.solanaceaesource.org).
Taxonomic synopsis of Solanum
endemic in the West Indies
Solanum acropterum Griseb., Fl. Brit. W.I.: 437. 1862
Type: Jamaica. sin. loc., J. Waters s.n. (lectotype, K
[K000005250], designated by Knapp, 2008b: 409).
Distribution. Endemic to Jamaica, where it is found
in woods on limestone soils in the cloud forests of the
John Crow Mountains from 100-700 m.
Representative specimen. Jamaica. St. Thomas
Parish, Stearn 499 (A, BM, K, MO, P, S).
Solanum bahamense L., Sp. Pl.: 118. 1753
Type: Bahamas. “Ins. Providentia”, Anon. (lectotype LINN 248.42 [BH neg. 6822], designated by
Knapp & Jarvis, 1990: 328).
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West Indian Solanum
Solanum racemosum Jacq., Enum. Syst. Pl.: 15. 1760
Type: Martinique, no specimens located (neotype,
Jacquin, Select. Stirp. Hist. Amer. t. 36. 1763, designated by Howard, 1989: 293).
Solanum igneum L., Sp. Pl., ed. 2: 160: 1762
Type: No specimens located (lectotype, “Solanum
spiniferum frutescens spinis igneis Americanum” in
Plukenet, Phytographia, t. 225, f. 5. 1692, designated
by Howard, 1989: 293).
Solanum fruticosum Mill., Gard. Dict. Ed. 8, no. 18.
1762
Type: Jamaica. sin. loc., W. Houstoun s.n. (lectotype,
BM [BM000815974, BH neg. 5109], designated here).
Solanum varginstonicum Buc’hoz, Pl. Nouv. Découv.
48, tab. 46. 1784
Type: Pl. Nouv. Découv. 48, tab. 46. 1784. Type:
Cultivated in Paris, no specimens located (lectotype,
Buch’hoz, Pl. Nouv. Decouv. 48, t. 46. 1784, designated here).
Solanum igneum L. var. parvifolium Vahl, Eclog.
Amer. 1: 23. 1797
Type: U.S. Virgin Islands. St. Croix, D. West s.n.
(holotype, C; isotype, G-DC [Morton neg. 8431]).
Solanum subarmatum Willd., Enum. Pl.: 240. 1809
Type: Cultivated at Berlin, of unstated origin, but
apparently thought to be from Bahamas “Insula Providentia”, Anon. (holotype, B-W [B-W-4414]).
Solanum persicifolium Dunal, Nat. Hist. Solanum
1813
Type: No specimens cited in protologue (lectotype,
Plumier, Pl. Amer. t. 244, fig. 2, 1760, designated
here). as “persicaefolium”.
Solanum anacanthum Dunal in DC., Prodr. 13(1):
183. 1852
Type: “Mexico”, J. Pavón s.n. [unspecified locality in
West Indies, M. Sessé & J. Mociño s.n.] (holotype, G
[G-Boiss., F neg. 34108]; isotype, MA [MA-604598]).
Solanum igneum L. var. inerme Dunal in DC., Prodr.
13(1): 184. 1852
Type: Cultivated, originally from St. Thomas, 1827,
H. Wydler 90 (lectotype, G-DC [IDC microfiche 80061.2077.III.8], designated here).
Solanum persicifolium Dunal var. angustifolium Dunal
in DC., Prodr. 13(1): 185. 1852
Type: “Puerto Rico, St. Thomas”, 1820, C. Bertero
s.n. (holotype, G-DC [F neg. 33949, IDC microfiche
800-61.2077.I.2]; isotype, AV-Requien n.v.).
Solanum bahamense L. var. lanceolatum Griseb., Fl.
Brit. W.I.: 440. 1861
69
Type: Jamaica.(“Solanum fruticosum bacciferum
spinosum, flore caeruleo”, lectotype, Sloane, Voy. Jamaica 1: t. 145, fig. 3, 1707, designated here; epitype,
BM-SL [BM000589942], designated here).
Solanum bahamense L. var. subarmatum (Willd.) O.E.
Schulz in Urb, Symb. Antill. 6: 222. 1909
Type: Based on Solanum subarmatum Willd.
Solanum racemosum Jacq. var. igneum (L.) O.E.
Schulz in Urb., Symb. Antill. 6: 225. 1909
Type: Based on Solanum igneum L.
Solanum persicifolium Dunal var. parvifolium (Vahl)
O.E. Schulz in Urb., Symb. Antill. 6: 227. 1909
Type: Based on Solanum igneum L. var. parvifolium
Vahl.
Solanum persicifolium Dunal var. belloi O.E. Schulz in
Urb., Symb. Antill. 6: 227. 1909
Type: Puerto Rico. along seashore near Fajardo, 17
Feb 1900, A.A. Heller 4605 (lectotype, F [F-80757],
designated here; isolectotypes, B [destroyed], MO,
NY [NY00144404], P [P00368434]).
Solanum bahamense L. var. luxurians D’Arcy, Ann.
Missouri Bot. Gard. 61: 838. 1974
Type: USA. Florida: Monroe Co., Key Largo, 21
Oct 1967, W.G. D’Arcy 2234 (holotype, FLAS; isotype, MO [MO-1963193]).
Solanum bahamense L. var. rugelii D’Arcy, Ann. Missouri Bot. Gard. 61: 839. 1974
Type: USA. Florida: Monroe Co., Key West, F.
Rugel s.n. (holotype, MO).
Distribution. Widespread in the West Indies in
coastal forests, forest margins and on beach margins,
often on coral or calcareous soils, from 0-100 m. Absent from Hispaniola, but found on Ile la Tortue and
Ile la Navasse off the NW coast of Haiti.
Representative specimen. Bahamas, Bimini, Cat
Cay, Correll & Correll 45600 (F, NY).
In describing Solanum fruticosum, Miller (1768)
cited “Solanum fruticosum bacciferum, spinosum, flore
caeruleo Sl. C. 108” and made reference to the collector William Houstoun. The specimen selected here as
the lectotype for this name in BM is annotated with
the same reference to Sloane, and bears a label in
Houstoun’s hand (Fig. 1a). It is also a good match for
the description.
Solanum varginstonicum was named by Buc’hoz in
honour of George Washington, with a spectacular
misspelling of the name. No specimens have been
traced for Buc’hoz’s names, so the plate (Fig. 1b) is selected here as the lectotype.
In his description of Solanum persicifolium, Dunal
Anales del Jardín Botánico de Madrid 66(1): 65-84, enero-junio 2009. ISSN: 0211-1322. doi: 10.3989/ajbm.2209
70
S. Knapp
(1813) cites two pre-Linnean sources; Plumier’s
(1703) catalogue (“Plumier cat. 4”) bound with the
Nova Plantarum Americanarum (Plumier, 1703) and
the larger compilation of Plumier’s American plants
illustrated by Johannes Burman from tracings of
Plumier’s originals held in Paris (“Burm., Plant.
Amer. p. 240, t. 244, fig. 2”; Plumier, 1755-1760, see
Jarvis, 2007 for a discussion of Plumier’s plates). On
page 4 in Plumier’s 1703 catalogue is the polynomial
“Solanum fruticosum, persicae foliis, aculeatum”; this
same polynomial is associated with plate 244 in the illustrated edition (Plumier, 1757). In the Prodromus
(Dunal, 1852), Dunal cites several specimens, stating
that the Burman plate was “mala”. The Burman plate,
however, is the only verifiable element associated with
the original description (although Dunal may have
seen the originals in Paris) and is thus here selected as
the lectotype (Fig. 1c).
a
b
c
d
e
f
Fig. 1. Solanum bahamense L. a, Lectotype of S. fruticosum Mill., Houstoun s.n. (BM000815974); b, lectotype of S. varginstonicum
Buc’hoz, Buc’hoz, Pl. Nouv. Découv. 48., t. 46, 1784. Reproduced with permission of the Natural History Museum Botany Library; c,
lectotype of Solanum persicifolium Dunal, Plumier, Plant. Amer. t. 245, f. 2, 1763. Reproduced with permission of the Natural History
Museum Botany Library; d, lectotype of S. bahamense var. lanceolatum Dunal, Sloane, Voy. Jamaica t. 145, fig. 3, 1707. Reproduced
with permission of the Natural History Museum Botany Library; e, epitype of S. bahamense var. lanceolatum Dunal (BM000589942).
Reproduced with permission of the Natural History Museum, London; f, lectotype of S. persicifolium var. belloi O.E. Schulz, Heller 4605
(F-80757). Reproduced with permission of the Field Museum of Natural History.
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West Indian Solanum
71
Solanum igneum var. inerme was described by
Dunal using two sheets in G-DC (Dunal, 1852), one
without a collector from Cuba and the other a cultivated specimen from seeds send by Wydler from St.
Thomas. He speculates that the un-armed nature of
the plant is due to it being cultivated (Dunal, 1852),
and places S. subarmatum tentatively in synonymy.
The only unambiguously identifiable sheet cited is
Wydler 90 from St. Thomas in the U.S. Virgin Islands;
it is here selected as the lectotype (not illustrated).
In describing Solanum bahamense var. lanceolatum,
Grisebach unambiguously cites a plate from Sloane as
part of the diagnosis, and additionally assigned some
of the specimens cited (without herbarium attribution) to this variety. I feel that he is specifically referring to the Sloane plate as the element on which he is
basing his name, and merely identifying the other collections as belonging to one or the other variety, thus
these specimens are probably not type material. Two
specimens at GOET annotated “? lanceolatum” in
Grisebach’s hand are potential original material, but
the Sloane plate is the only element unambiguously
linked to the variety. I therefore have chosen the plate
in Sloane’s A voyage to the islands Madera … Jamaica
(1707-1725) labelled Solanum fruticosum bacciferum
spinosum, flore caeruleo” as the lectotype of the variety (Fig. 1d). The epitype of this name with the same
polynomial designation is found in the Sloane herbarium at BM (BM000589942, Fig. 1e).
Five collections were cited in the protologue of
Solanum persicifolium var. belloi; Heller 4605, Sintenis
1260, Sintenis 1919 p.p., Eggers 35 and Benzoin s.n.
Of these, Heller 4605 is the most widely distributed
and has several sheets in very good condition and the
sheet at F (Fig. 1f) is here selected as the lectotype.
Representative specimen. Cuba, Pinar del Río, Ekman 17544 (S).
Solanum boldoense has occasionally been confused
with the Mexican and Central American species S.
dulcamaroides Dunal, to which it is probably closely
related.
Solanum boldoense Dunal & A.DC., in Prodr. 13(1):
679. 1852
Type: Based on Solanum cardiophyllum Dunal, non
Lindl.
Solanum cardiophyllum Dunal in DC., Prodr. 13(1)
89. 1852. Cuba. “in Havana”, B.M. Boldo s.n. (holotype, MPU).
Solanum scandens Sessé & Moc., Fl. Mex., ed. 2, 53.
1894
Type: Cuba. Sessé & Mociño s.n. (lectotype, MA
[MA-604676, F neg. 48318], designated by Knapp,
2008a: 20; isolectotype MA [MA-604675, F neg.
48343, 5381, upper frag. in photo, sheet now divided]).
Type: US Virgin Islands. St. John: Coral Bay, L.C.
Richard s.n. (holotype, P [P00325582]; isotype fragment, F [F-976721]).
Distribution. Endemic to Cuba, where it is found in
forests and forest edges at low to middle elevations.
Solanum chamaecanthum Griseb., Cat. Pl. Cub.: 190.
1866
Type: Cuba. “in savanis prope Hanabana”, 1865, C.
Wright s.n. (lectotype, GOET [GOET0001970], designated here; isolectotype, GOET [GOET0001971];
possible isolectotypes (numbered 3634), GH [GH77546], K [K000196224], NY [NY00111375,
NY00689170], S [S04-2892]).
Distribution. Endemic to Cuba, in low elevation
swampy areas, common in the northwestern part of
the island around Havana.
Representative specimen. Cuba, Camagüey, Ekman
8600 (S).
In describing S. chamaeacanthum, Grisebach
(1866) did not cite a specific Wright collection number. Wright’s duplicates are not all numbered, and
some distributed without numbers are clearly duplicates of numbered collections. A sheet at GOET
(GOET0001970) is labelled “savanas Habanana” and
is an un-numbered Wright collection, making it the
clear choice for a lectotype (Fig. 2a). Another sheet
(GOET0001971) was annotated lectotype by R.A.
Howard in 1984, but as it does not have the locality associated with the protologue has not been selected
here as the lectotype.
Solanum conocarpum Rich. ex Dunal, in Poir., Encycl. Suppl. 3: 748. 1814
Distribution. Endemic to the U.S. Virgin Islands,
found in coastal thickets.
Representative specimen. U.S. Virgin Islands, St.
John, Acevedo 5437 (K, MO).
Solanum crotonoides Lam., Tabl. Encycl. 2: 24. 1794
Type: “Martinique”. sin. loc., J. Martin s.n. (holotype, P-LA [P00357672]).
Solanum reticulatum Dunal in Poir., Encycl. Suppl. 3:
764. 1814
Type: “Saint Domingue”, [Hispaniola], P. Richard
s.n. (holotype, P [P00325752, Morton neg. 8176];
fragment, F).
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S. Knapp
a
b
c
d
Fig. 2. a, lectotype of Solanum chamaeacanthum Griseb., Wright s.n. (GOET0001971); b, lectotype of S. hoplophorum O.E. Schulz,
Fuertes 1023 (S04-2916) (= S. crotonoides Lam.). Reproduced with permission of the Swedish Museum of Natural History; c, lectotype
of S. formonense O.E. Schulz and of S. formonense var. parvifolium O.E. Schulz, Ekman H.10265 (S04-2911) (= S. crotonoides Lam.).
Reproduced with permission of the Swedish Museum of Natural History; d, lectotype of S. dendroicum O.E.Schulz & Ekman, Ekman
H.15102 (S04-2902). Reproduced with permission of the Swedish Museum of Natural History.
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West Indian Solanum
Solanum hoplophorum O.E. Schulz in Urb., Symb.
Antill. 7: 369. 1912
Type: Dominican Republic. Barahona, Gippel des
Noche Buena, 1800 m, Aug 1911, M. Fuertes 1023
(lectotype, S [S04-2916], designated here; isolectotypes, A [A00077555], BM [BM000629234], GH
[GH00077554], GOET, K [K000196486], MO
[MO-2572037]).
Solanum formonense O.E. Schulz ex O. Schmidt,
Repert. Spec. Nov. Regni Veg. 32: 92. 1933
Type: Haiti. Massif de la Hotte, in parte occidentali, Les Anglais, Morne l’Etang, 1400 m, 21 Jul 1928,
H. Ekman H.10365 (lectotype, S [S04-2911], designated here).
Solanum formonense O.E. Schulz ex O. Schmidt var.
grandifolium O.E. Schulz ex O. Schmidt, Repert.
Spec. Nov. Regni Veg. 32: 92. 1933
Type: Haiti. Massif de la Hotte, western group,
Torbec, M. Formon, 1700 m, 30 Dec 1926, E.L. Ekman H.7461 (lectotype, S, designated here).
Solanum formonense O.E. Schulz ex O. Schmidt var.
parvifolium O.E. Schulz ex O. Schmidt, Repert.
Spec. Nov. Regni Veg. 32: 92. 1933
Type: Haiti. Massif de la Hotte, western group, Les
Anglais, Morne l’Etang, 1400 m, 21 Jul 1928, H. Ekman
H.10365 (lectotype, S [S04-2911], designated here).
Distribution. Known only from the montane cloud
forests of Hispaniola (Haiti and the Dominican Republic), from 1000-1800 m elevation.
Representative specimen. Dominican Republic. Prov.
Peravia, Zanoni & Pimentel 40315 (JBSD, MO, NY).
Although Lamarck’s protologue indicates that the
plant collected by Joseph Martin came from Martinique, Solanum crotonoides has never been collected there. A similar situation exists for S. pyrifolium (Dulcamaroid clade), also collected by Martin
from “Martinique”. I suspect that Martin did in
fact also visit Hispaniola (the locality for his type collection of Aquartia microphylla is “S. Domingo”, see
below) or that Martin used Martinique as a broad
locality covering most of the Caribbean region, and
that both these plants are true Hispaniolan endemics.
In describing S. formonense, Schulz and Schmidt
did not designate which of the two varieties was typical. Sheets of both syntypes are held at S; Ekman
H.10365 (S04-2911, Fig. 2b) is in excellent condition
and is here used to lectotypify both the species and the
variety. Ekman H.7461 is the lectotype for var. grandifolium, but is not figured here.
The lectotype of S. hoplophorum (Fig. 2c) is a very
robust, prickly male plant (Fuertes 1023), showing
73
that combination of characters that Dunal (1814) noted in describing S. reticulatum as the “hermaphrodite” of S. crotonoides (which is a dioecious species,
see Knapp & al., 1998). Both varieties of S. formonense are female plants.
Solanum dendroicum O.E. Schulz & Ekman, Repert.
Spec. Nov. Regni Veg. 32: 91. 1933
Type: Dominican Republic. Prov. Samaná, Peninsula de Samaná, near Samaná, Laguna, ca. 350 m, 21
May 1930, E.L. Ekman H.15102 (lectotype, S [S042902], designated here; isolectotypes, A [A0077457],
K [K000196485], NY [NY00111365]).
Distribution. Endemic to the Samaná Peninsula of
the Dominican Republic in northeastern Hispaniola,
in matorrales and forests on limestone derived from
ancient corals from sea level to 350 m elevation.
Representative specimen. Dominican Republic,
Prov. Samaná, Peguero & Veloz 549 (JBSD, MO).
Of the four syntypes cited by Schulz (Ekman
H.14847, Ekman H.15042, Ekman H.15102 and Abbott 2922), only one (Ekman H15102) is fertile. The
sheet in S (Fig. 2d) has been chosen as the lectotype
because Eric Ekman’s top set of specimens is housed
in Stockholm, all have been annotated by Ekman, and
S04-2902 is in excellent condition.
Solanum ensifolium Dunal, in DC., Prodr. 13(1): 186.
1852
Type: “Mexico, J. Pavón” [Puerto Rico, M. Sessé &
J. Mociño s.n.] (holotype, G [G-Boiss., F neg. 34117];
isotype, MA [MA-604619, F neg. 48338]).
Solanum drymophilum O.E. Schulz in Urb., Symb.
Antill. 6: 227. 1909
Type: Puerto Rico, Cayey, Monte Llano, 25 Sep
1885, P. Sintenis 2374 (lectotype, BM [BM000886627],
designated here; isolectotypes A [A00242857], CORD,
F [F-79600, F-680330], GH [GH00242855], GOET,
K [K000438174], M, MO [MO-2792753, MO5467986, MO-5468002], NY [NY00111367], P
[P00367425, P00367426, P00367427, P00367428,
P00367429], S, W).
Distribution. Endemic to Puerto Rico, in forests on
low hills in the southern part of the island, to 800 m.
Representative specimen. Puerto Rico, Mun. Salinas, Proctor 41918 (NY).
Many syntypes were cited in the original description of S. drymophilum; Sintenis 2025, Sintenis 2217,
Sintenis 2121, Sintenis 2374, Sintenis 2480, Sintenis
5401. Of these, Sintenis 2374 is represented in many
herbaria, and is composed a number of sheets in good
condition with both flowers and fruit. I have selected
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S. Knapp
the BM sheet as the lectotype (Fig. 3a) because it has
all reproductive stages and is in good condition.
Solanum gundlachii Urb., Symb. Antill. 5: 487. 1908
Type: Cuba. sin. loc., C. Wright 3030 (lectotype,
GH [GH00077553], designated here; isolectotypes,
BM [BM000886657], MO [MO-2766697], GOET [2
sheets], K [K000196481], MA [MA-607704]).
Solanum hotteanum Urb. & Ekman, Ark. Bot. 22(10):
51. 1929
Type: Haiti. Massif de la Hotte, western group, Tarbec, top of Morne Formon, ca. 2225 m, 1 Jan 1927,
E.L. Ekman H.7489 (holotype, S [S04-2917]).
Solanum gundlachii Urb. var. monteverdense O.
Schulz, in Urban, Symb. Antill. 6: 210. 1909
Type: Cuba. Monteverde, 800 m, H.F.A. von Eggers
5108 (holotype, B destroyed, no duplicates yet found).
Solanum lomense Britton & P. Wilson, Bull. Torrey
Bot. Club 53: 464. 1926
Type: Cuba. Prov. Oriente, Loma San Juan, near
Loma del Gato, Hongolosongo, 22 Apr 1926, Bro.
Clement 2520 (holotype, NY [NY00111364]) (“Bro.
León” as collector in protologue). as “lomensis”.
Solanum cristalense Amshoff, Rev. Soc. Cubana Bot.
13: 6. 1956
Type: Cuba. Oriente: Pinares, el Prado, Nicaro, Mayari, J. Acuña & F. Zaguas 19778 (holotype, SV n.v.).
Solanum cristalense Amshoff var. macrospermum
Hada?, Folio Geobot. Phytotax. Prague 5: 431. 1971
Type: Cuba. Oriente: montibus Montecristo, E.
Hada? 1421 (holotype, P).
Distribution. Only known from eastern Cuba and
from Haiti, in pine woods from 700-2225 m elevation.
Representative specimen. Cuba, Santiago de Cuba,
Herbario Flora de Cuba (HFC) 57223 (B).
The type collection of Solanum gundlachii (Wright
3030) is present in many herbaria, some of which
have both flowers and fruit. The specimen at GH
[GH00077553] has both flowers and mature fruits
and is here selected as the lectotype. The sheet clearly
represents two separate gatherings, as two labels bear
different collection dates and appear to come from
different localities (see Fig. 3b).
Solanum havanense Jacq., Enum. Syst. Pl. 15. 1760
Type: Jamaica. Tab. 35, Jacq., 1763 (neotype, tab.
35 in Jacquin, Select. Stirp. Amer. Hist. 1763, designated by Knapp, 2008b: 414).
Solanum coriaceum Hook., Bot. Mag. 54: tab. 2708.
1827
Type: Cultivated in England, the seeds said to be
sent by R. Barclay from Mexico (lectotype, Hooker,
Bot. Mag. 54: t. 2708. 1827, designated by Knapp,
2008b: 414).
Solanum hookerianum A. Spreng., Tent. Suppl. Syst.
9. 1828
Type: Based on Solanum coriaceum Hook.
Solanum myrtifolium Lodd., Bot. Cab. no. 1431. 1828
Type: Cultivated in London (lectotype, Loddiges,
Bot. Cab. 25: t. 1431. 1828, designated by Knapp,
2008b: 414).
Solanum havanense Vell., Fl. Flumin. 82. 1829; Icones
2, tab. 94. 1831
Type: Icon. Tab. 94, Brazil. Rio de Janeiro: Pharmocopolis (lectotype, Vellozo, Fl. Flumin. Icones 2:
t. 94. 1831, designated by Knapp, 2008b: 414).
Diamonon coriaceum (Hook.) Raf., Fl. Tellur. 2: 76.
1836
Type: Based on Solanum coriaceum Hook.
Solanum byrsinum Voigt, Hort. Suburb. Calc. 511.
1845
Type: Based on Solanum coriaceum Hook.
Solanum vacciniifolium Dunal, in DC., Prodr. 13(1):
160. 1852
Type: Cuba. Havana, R. de la Sagra 294 (holotype, GDC [F neg. 33944, IDC microfiche 800-61.2075:I.2]).
Solanum havanense Jacq. var. vacciniifolium (Dunal)
Griseb., Cat. Pl. Cub. 189. 1866
Type: Based on Solanum vacciniifolium Dunal.
Distribution. Endemic to the West Indies, found on
Cuba, Jamaica and the Cayman Islands, in coastal
woodlands on calcareous soils.
Representative specimen. Cuba, Cienfuegos, Webster & al. 51 (A).
Solanum maestrense Urb., Repert. Spec. Nov. Regni
Veg. 21: 221. 1925
Type: Cuba. Oriente: Sierra Maestra, on the water
divide between Rio Yara and Río Palmamocha, at the
foot of Loma Joaquim, on the slope towards Rio Palmamocha, 19 Jul 1922, E. Ekman 14456 (lectotype, S
[S04-2941], designated here; isolectotype, S [S042942]).
Distribution. Endemic to Cuba, known only from
the type.
Representative specimen. Cuba, Oriente, Ekman
14456 (S).
It is clear from the protologue that Urban did not
have at his disposal a specimen with a flower (“flores
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West Indian Solanum
75
a
b
c
d
Fig. 3. a, lectotype of Solanum drymophilum O.E Schulz, Sintenis 2374 (BM000886627) (= S. ensifolium Dunal). Reproduced with permission of the Natural History Museum, London; b, lectoytpe of S. gundlachii O.E. Schulz, Wright 3030 (GH00077553). Reproduced
with permission of Harvard University, Cambridge, Massachusetts, USA; c, lectotype of S. maestrense Urb., Ekman 14456 (S04-2941).
Reproduced with permission of the Swedish Museum of Natural History; d, lectotype of S. squarrosum O.E. Schulz, Schomburgk 51
(L0403808). Reproduced with permission of the Natural History Museum of the Netherlands.
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S. Knapp
non vidi”, Urban, 1925), and that he was describing
this species from material in B that is no longer extant.
The two rather scrappy sheets at S are both annotated
“type”, but only one (S04-2941) has reproductive
parts (one flower and two fruits in the packet). This
sheet (Fig. 3c) is chosen here as the lectotype.
Solanum microphyllum (Lam.) Dunal, Hist. Solan.
187. 1813
Type: Based on Aquartia microphylla Lam.
Aquartia microphylla Lam., Tabl. Encycl. Meth. 1:
327, tab. 82, fig. 2. 1791
Type: “S. Domingo” [Dominican Republic], J.
Martin s.n. (holotype, P-LA [P00357664]).
Solanum microphyllum (Lam.) Dunal var. tomentosum Dunal in DC., Prodr. 13(1): 193. 1852
Type: “Santo Domingo”, C. Bertero s.n. (holotype
G-DC [IDC microfiche 800-61.2077:II.6]; isotype
(probable), MO [MO-5468004]).
Solanum didymacanthum Millsp., Publ. Field
Columbian Mus., Bot. Ser. 2: 183. 1906
Type: Bahamas. Exuma: Hayne’s road, 22-28 Feb
1905, N. Britton & C. Millspaugh 3037 (holotype, F [F174174, F neg. 51388]; isotypes, K [K, NY [NY
00111366], US [US-849507, barcode US00027549]).
Solanum squarrosum O.E. Schulz in Urb., Symb. Antill. 6: 197. 1909
Type: Dominican Republic. “prope Azua”, R. Schomburgk 51 (lectotype, L [L0403808], designated here).
Solanum gonaivense Leonard, J. Wash. Acad. Sci. 17:
71. 1927
Type: Haiti. Dept. Artibonite, 8 mi NE of Gonaives, Ennery Road, 19 Feb 1926, E. Leonard 10007
(holotype, US [US-1301034, barcode US00588712];
isotype, GH [GH0077550]).
Distribution. In dry forests and xerophytic scrublands on Hispaniola and the Bahamas, usually on
limestone soils at around sea level, but up to 200 m elevation.
Representative specimen. Bahamas, Little Exuma,
Correll & Wasshausen 51203 (MO, NY, US).
The type specimen of Solanum squarrosum was cited as Schomburgk 51 and was at B, but is no longer extant. A rather scrappy duplicate of Schomburgk 51 at
Leiden (L0403808, Fig. 3d) is labelled “typus” in
Schulz’s hand and is here selected as the lectotype.
Solanum moense Britton & Wilson, Mem. Torr. Bot.
Club 16: 101. 1920
Type: Cuba. Oriente: Camp La Gloria, south of
Sierra Moa, 24-30 Dec 1910, J.A. Schafer 8125 (holo-
type, NY [NY00111363]; isotype, US [US-01047827,
barcode US00027693]).
Distribution. Known only from a few collections in
the Sierra de Moa in the eastern mountain ranges of
Cuba.
Representative specimen. Cuba, Holguin, Herbario
Flora de Cuba (HFC) 56400 (B).
Solanum orthacanthum O.E. Schulz in Urb., Symb.
Antill. 7: 537. 1913
Type: Dominican Republic. Azua prov.,Valle de
Yaque del Sur, 1300 m, Aug, M. Fuertes 1890 (lectotype, A [A00077559], designated here).
Solanum coelocalyx Alain, Phytologia 22: 170. 1971
Type: Dominican Republic. Constanza, El Motanzo,
A.H. Liogier & E.J. Marcano 14716 (holotype, NY
[NY00111376]; isotypes, BM [BM000886620], F [F1773526], GH [GH00231758]).
Distribution. Endemic to the eastern part of the island of Hispaniola in the Dominican Republic in
moist areas in montane pine forest from 1200-1400 m
elevation.
Representative specimen. Dominican Republic.
Prov. La Vega, Zanoni & al. 23215 (JBSD).
The sheet of Fuertes 1890 at A (Fig. 4a) is the only
duplicate of the type collection I have been able to
trace, and thus I have chosen it as the lectotype of
Solanum orthacanthum.
Solanum pachyneuroides Amshoff, Acta Bot. Neerland. 3: 418. 1954
Type: Cuba. Road to Delta Mine, Punta Gorda,
Moa region, Jul 1949, Bro. Alain, Bro. Clemente &
Bro. Chrysogone 912 (holotype, LS n.v.; isotype, US
[US-2815217, barcode US00027721]).
Solanum pachyneurum O.E. Schulz var. glabrescens
O.E. Schulz in Urb., Symb. Antill. 6: 196. 1909
Type: Cuba. Sin. loc., 1860-1864, C. Wright 382
p.p. (holotype, B, destroyed; lectotype, GH
[GH00112906, left hand stems, see below], designated here; isolectotype, K [K000196233]).
Distribution. Endemic to Cuba, known only from
the eastern part of the island from pine woods at ca.
700 m elevation.
Representative specimen. Cuba, Guantanamo,
Herbario Flora de Cuba (HFC) 42589 (B).
Wright 382, the type collection of Solanum
pachyneurum var. glabrescens and a syntype of S.
pachyneurum, is a mixed collection of S. pachyneurum
(e.g., the sheet at BM [BM000886750] is S. pachyneurum) and S. pachyneuroides (also see below). I have
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West Indian Solanum
77
a
b
c
d
Fig. 4. a, lectotype of Solanum orthacanthum O.E. Schulz, Fuertes 1890 (A00077559). Reproduced with permission of Harvard University, Cambridge, Massachusetts, USA; b, lectotype of S. pachyneurum O.E. Schulz var. glabrescens O.E. Schulz, Wright 382 p.p. (left
hand stems of GH00112906) (= S. pachyneuroides). Reproduced with permission of Harvard University, Cambridge, Massachusetts,
USA; c, lectotype of S. pachyneurum O.E. Schulz, Linden 1944 (K000196232). Reproduced with permission of the Royal Botanic Gardens, Kew; d, lectotype of S. polygamum Vahl var. thomae O. Kuntze, Kuntze 36 (NY0074477) (= S. polygamum). Reproduced with
permission of the New York Botanical Garden.
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S. Knapp
chosen the left-hand two stems of GH00112906 (Fig.
4b) to lectotypify the varietal name; the right-hand
stem and a small stem in the lower left corner are S.
pachyneurum.
Solanum pachyneurum O.E. Schulz in Urb., Symb.
Antill. 6: 195. 1909
Type: Cuba. Guantánamo: Santiago, prope Monte
Libanon [Monte Libano], 1843, J. Linden 1944 (lectotype, K [K000196232], designated here; isolectotype,
BM [BM000886751]).
Distribution. Endemic to eastern Cuba, in forests at
about 700 m elevation.
Representative specimen. Cuba, Guantanamo, Herbario Flora de Cuba (HFC) 52625 (B).
Schulz (1909) cited three collections in his original
description of Solanum pachyneurum – Wright 382,
Eggers 5197 and Linden 1944. Wright 382, original
material of both Solanum pachyneurum and S. pachyneurum var. glabrescens (a synonym of S. pachyneuroides, see above), consists of a series of mixed sheets
of S. pachyneurum and S. pachyneuroides. It is not
clear whether the two taxa occur sympatrically, or
whether Wright assigned his numbers to plants collected after the fact, common practice at the time (and
very likely). The latter case is suggested by the labeling
of the sheet of Wright 382 at GH (see Fig. 4b). The
sheet that Schulz examined in B has been destroyed,
and other sheets of Wright 382 are mostly of S.
pachyneurum, but some are a mixture of S. pachyneurum and S. pachyneuroides (see above). For this reason, it is more sensible to choose one of the many other syntypes of S. pachyneurum as the lectotype to avoid
confusion. A sheet of Linden 1944 was not found at B
and is presumably destroyed, but well preserved duplicates have been located at BM and K. Neither the K
nor the BM sheet has any locality data, but the sheet at
K (Fig. 4c) is better preserved, has reproductive parts
and has therefore been selected as the lectotype.
Solanum polyacanthos Lam., Tabl. Encycl. 2: 23. 1794
Type: Cultivated in Paris from seeds from “St.
Domingue” (Hispaniola), Anon. s.n. (holotype, P-LA
[P00357671]).
Solanum parviflorum Cav., Icon. 3: 19, tab. 236. 1795
Type: Cultivated in Paris, from Santo Domingo,
1786, Anon. s.n. (lectotype, MA [MA-206020], designated by Knapp, 2007: 198).
Solanum polyacanthos Lam. var. oreophilum O.E.
Schulz in Urb., Symb. Antill. 6: 200. 1909
Type: Haiti. Sin. loc., Nectoux s.n.; Furcy, Morne
Tranchant, 1800 m, L. Picarda 245; sin. loc., Christ
1757; Kenshoff, 1400 m, W. Buch 915; Gonaives, Furcy, 1500 m, Oct 1903, W. Buch 949. as “polyacanthos”.
Solanum tortipes Urb. & Ekman, Ark. Bot. 21(5): 59.
1927
Type: Haiti. Massif des Cahos, Petit Rivière de l’Artibonite, Pérodin, Jardins Nan-Plaine, 1400 m, 8 Mar
1925, E.L. Ekman H.3475 (holotype, S [S-04-2991];
isotype, US [US-1410723, barcode US00028728]).
Distribution. Known only from the island of Hispaniola (Haiti and the Dominican Republic), growing
in a wide variety of forest types from dry thorn forest
to cloud forest, from 300-1900 m.
Representative specimen. Dominican Republic,
Prov. San Juan, Howard & Howard 8805 (BM).
I have located no duplicates of the five specimens
were cited by Urban in describing Solanum polyacanthos var. oreophilum – Nectoux s.n., Picarda 245, Buch
915 and Buch 949, all from Haiti. A photograph of a
duplicate of Buch 949 at IJ is held at MO and is clearly S. polyacanthos. I have not lectotypified this name
here in the hopes of eventually finding and personally
examining duplicates of the syntype material.
Solanum polygamum Vahl, Symb. Bot. 3: 39. 1794
Type: U.S. Virgin Islands. St. Croix, H. West s.n.
(holotype, C).
Solanum inclusum Grisebach var. albiflorum Eggers,
Bull. U.S. Natl. Mus. 13: 77. 1879 [also included in
Smithson. Misc. Colls. 23(3): 77. 1882]
Type: No specimen or specific locality cited.
Solanum polygamum Vahl var. thomae Kuntze, Revis.
Gen. Pl. 2: 455. 1891
Type: U.S. Virgin Islands. Saint Thomas: sin. loc.,
O. Kuntze 36 (lectotype NY [NY0074477], designated here; isolectotype, K [K000196236]).
Distribution. Found in the central Caribbean on St.
Thomas, St. Jan and St. John (US Virgin Islands), Tortola and Virgen Gorda (British Virgin Islands), Isla
Vieques just to the southeast of Puerto Rico, in eastern
Cuba and on southern Hispaniola from 0-350 m in
limestone and sandy areas.
Representative specimen. British Virgin Islands,
Virgin Gorda, Little & al. 23829 (BM).
Two syntypes were cited by Kuntze for Solanum
polygamum var. thomae; Eggers 100 and Kuntze 36.
Sheets of both of these were located at K; but the duplicate of Kuntze 36 at NY (Fig. 4d) is annotated by
Kuntze and comes from his personal herbarium and is
thus selected as the lectotype. Solanum inclusum var.
albiflorum was described by Eggers in his Flora of St.
Croix and the Virgin Islands (1879) without citing a
Anales del Jardín Botánico de Madrid 66(1): 65-84, enero-junio 2009. ISSN: 0211-1322. doi: 10.3989/ajbm.2209
West Indian Solanum
specimen or a specific locality, but is clearly a synonym of S. polygamum. The description is of a female
plant (S. polygamum is dioecious, see Knapp & al.,
1998), and I prefer to continue to search for fruiting
material annotated by Eggers S. polygamum before
neotypifying this name.
Solanum punctulatum Dunal in DC., Prodr. 13(1):
122. 1852
Type: Jamaica. sin. loc., 1819, J. Wiles [in herb.
Lambert] s.n. (holotype, G [Morton neg. 8638]).
Distribution. Endemic to Jamaica, where it is found
in forests and forest edges at high elevations.
Representative specimen. Jamaica, St. Thomas
Parish, Stearn 99 (BM, K, S).
Solanum pyrifolium Lam., Tabl. Encycl. 2: 19. 1794
Type: “Martinica”, sin. loc. [?Hispaniola], J. Martin s.n. (holotype, P-LA [P00357682]).
Solanum domingense Dunal in DC., Prodr. 13(1): 80.
1852
Type: Dominican Republic, sin. loc., Anon. s.n.
(holotype, G [F neg. 33943]).
Distribution. Known only from the island of Hispaniola, primarily in the Dominican Republic, along
forest edges on limestone and in cloud forests, from
120-1600 m elevation.
Representative specimen. Haiti, Holdridge 935
(BM, F).
Solanum schulzianum Urb., Symb. Ant. 5: 488. 1908
Type: Dominican Republic. Jimenes, 1190 m, 31
May 1887, H.F.A. von Eggers 2208 (lectotype, K
[K000196492], designated here).
Solanum abbottii Leonard, J. Wash. Acad. Sci. 14:
415. 1924
Type: Dominican Republic. Prov. Pacificador,
Quita Espuela, vic. San Francisco de Macoris, 900 m,
7 Apr 1922, W. Abbott 2073 (holotype, US [US1079509, barcode US0027429]; isotype, NY
[NY00111371]).
Distribution. Known only from the eastern portion
of the island of Hispaniola (collections only seen from
the Dominican Republic) growing in forests from
300-1300 m elevation.
Representative specimen. Dominican Republic,
Prov. Samana, Salazar & al. 1309 (JBSD).
Of the two syntypes cited in the original description (Eggers 1608, Eggers 2208), only Eggers 2208 was
cited by Urban as being fertile; the Kew duplicate
79
(Fig. 5a) of this collection is therefore the most appropriate choice for a lectotype.
Solanum selleanum Urb. & Ekman, Ark. Bot. 21(5):
60. 1927
Type: Haiti. Massif de la Selle, Morne de la Selle,
Croix des Bouquets, above Badeau, 1600 m, 28 Jan
1925, E.L. Ekman H.3127 (lectotype, S [S04-2981],
designated here; isolectotypes, S [S-04-2982], US
[US-1304052, barcode US00027793]).
Distribution. Only known from the island of Hispaniola, in valleys (Morne de la Selle, Haiti) at ca.
1600 m. The type and one other sheet from the Dominican Republic (Zanoni & al. 33773, not seen) are
the only specimens of this species.
Representative specimen. Haiti, Ekman H.3127 (S).
Two sheets of Ekman H.1327 are in S, both annotated by Ekman. Of these, S04-2981 (Fig. 5b) is more
complete, with both flowers and fruit, and is here selected as the lectotype.
Solanum tetramerum Dunal in DC., Prodr. 13(1):
681. 1852
Type: Cuba. Guantánamo, J. Linden 1986 (holotype, G-DC; isotypes, BM [BM000886483], K
[K000196465], MO [MO-1058522]).
Aquartia aculeata Jacq., Enum. Pl. Carib. 1: 12. 1760
Type: Dominican Republic. “Domingo, ad sinum
Bayaha” (no specimen located; neotype, Jacquin, Select. Stirp. Hist. Amer. t. 12. 1763, designated here).
Solanum aquartia Dunal, Hist. Solan. 187. 1813
Type: Based on Aquartia aculeata Jacq. Nom. illeg.
Solanum inclusum Grisebach, Fl. Brit. W. Ind. 441.
1861
Type: Cuba. sin loc., J. Linden 1986 (neotype, BM
[BM000886483], designated here; isoneotypes, K
[K000196465], G-DC, MO [MO-1058522]).
Solanum grisebachianum Kuntze, Revis. Gen. Pl. 2:
455. 1891
Type: Based on Solanum inclusum Grisebach.
Solanum cahosianum Urb. & Ekman, Ark. Bot.
22(10): 49. 1929
Type: Haiti. Massif des Cahos, near St.-Michel,
road between Pol and Dessalines, 450 m, 8 Jun 1927,
E.L. Ekman H.8378 (holotype, S [S04-2889]).
Solanum miragonae Urb., Ark. Bot. 22(10): 50. 1929
Type: Haiti. Massif de la Hotte, group Morne Rochelois, Miragoane, limestone cliffs towards Anse-àVeau, 23 Nov 1926, E.L. Ekman H.7289 (lectotype, S
[S04-2952]).
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80
S. Knapp
a
b
d
c
Fig. 5. a, lectotype of Solanum schulzianum Urb., Eggers 2208 (K000196492); b, lectotype of S. selleanum Urb. & Ekman, Ekman
H.3127 (S04-2981). Reproduced with permission of the Swedish Museum of Natural History; c, lectotype of Aquartia aculeata Jacq.,
Jacquin, Select. Stirp. Amer. t. 12, 1763) (= S. tetramerum Dunal). Reproduced with permission of the Natural History Museum Botany
Library; d, neotype of S. inclusum Griseb., Linden 1986 (BM000886483) (= S. tetramerum Dunal). Reproduced with permission of the
Natural History Museum, London.
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West Indian Solanum
Solanum aculeatum (Jacq.) O.E. Schulz in Urban,
Symb. Ant. 6: 196. 1909
Type: Based on Aquartia aculeata Jacq.
Solanum aculeatum (Jacq.) O.E. Schulz var. fuertesii
O.E. Schulz in Urban, Symb. Ant. 7: 370. 1912
Type: Dominican Republic. Barahona prov., “am
Duvergé fluss”, 50 m, Aug 1910, M. Fuertes 384 (lectotype, NY [NY00111372], designated here; isolectotypes, E [E00202467], US [US-658349, barcode
US00027440]).
Solanum aculeatum (Jacq.) O.E. Schulz var. luxurians
O.E. Schulz ex O. Schmidt, Repert. Spec. Nov.
Regni Veg. 32: 91. 1933
Type: Haiti. Massif des Matheux, Thomazeau,
Morne-a-Cabrits, 400 m, 17 May 1928, E.L. Ekman
H.9979 (lectotype, S [S04-2827], designated here; isotype, K [K000196473]).
Solanum aquartia Dunal var. fuertesii (O.E. Schulz)
Alain, Phytologia 70: 150. 1991
Type: Based on Solanum aculeatum (Jacq.) O.E.
Schulz var. fuertesii O.E. Schulz.
Solanum aquartia Dunal var. luxurians (O.E. Schulz)
Alain, Phytologia 70: 150. 1991
Type: Solanum aculeatum (Jacq.) O.E.Schulz var.
luxurians O.E. Schulz ex O. Shmidt.
Distribution. In forests and scrub on limestone dry
hills on Cuba, Hispaniola and Jamaica, from sea level
to 400 m elevation.
Representative specimen. Jamaica, Saint Catherine
Parish, Proctor 32768 (BM).
There has long been confusion over the correct
name for this species. Dunal’s (1813) re-naming of
Jacquin’s Aquartia aculeata as Solanum aquartia was illegitimate, but the name was used for many years. By
the time Schulz (1909) made the correct combination
Solanum aculeatum, the name Solanum aculeatum St.Lag. had been validly (but illegitimately) published as
a replacement for Solanum aculeiger Dunal (now
recognised as a synonym of the Brazilian species S.
thomasiifolium Sendtn., see Solanaceae Source,
http://www.solanaceaesource.org). The oldest available name for this species is S. tetramerum.
No specimen of Aquartia aculeata attributable to
Jacquin has been found, despite intensive searches in
many herbaria (principally BM, LINN and W) where
such a sheet, if it had ever existed, might have been preserved. No original material was cited in the protologue
(Jacquin, 1760), so any type selected must be a neotype.
The illustration of A. aculeata in Jacquin’s first illustrated edition of the Selectarum Stirpium Americanarum
Historia (Jaqcuin, 1763), a copper engraving taken
81
from original drawings done by Jacquin in the field (see
Wiltshear, 1913) is very accurate, and I have chosen it
here as the neotype of the species (Fig. 5c).
Solanum inclusum was described without specific
collection or locality but said to be widespread across
the Caribbean and in northern South America (Grisebach, 1861). Various subsequent authors have suggested that Grisebach’s S. inclusum included both S.
tetramerum and S. polygamum. No specimens have
been identified at GOET that can be directly linked to
Grisebach’s naming of S. inclusum. In his Cuban catalogue Grisebach (1866) cited specimens as S. inclusum that are now identified as S. tetramerum (Linden 1986, 2000). Specimens indicated as type material of S. inclusum in some herbaria (i.e., Wright 1350 in
GH) are in fact S. torvum. Since Grisebach (1861) did
not indicate any specimens or specific localities for his
name, these sheets cannot be considered type material. I neotypify Grisebach’s Solanum inclusum here
with the BM duplicate (Fig. 5d) of Linden 1986 in order to stabilise usage (synonymy) of this ambiguous
name.
No duplicates of Ekman H.7289, the type of
Solanum miragonae, are extant at B, and the only sheet
of this collection encountered is at S, so this is the
clear choice as the lectotype (Fig. 6a). It may be that
the S sheet is the holotype, as S. miragonae was described in a Swedish journal along with other taxa
whose types are in S, but as there is no evidence of
this, I prefer to lectotypify the name here.
Solanum aculeatum var. fuertesii was described using several collections: Fuertes 384, 386 and 396. Of
these, Fuertes 384 has been located in several
herbaria, the NY duplicate is here selected as the lectotype (Fig. 6b). In describing variety luxurians, with
larger, more repand leaves, several syntypes were cited: Ekman H.2318, Ekman H.6732, Ekman H.8548,
Ekman H.9979 and Ekman H.8773. The duplicate of
Ekman H.9979 in S (Fig. 6c) has an annotation label
in Schulz’s hand with the varietal name luxurians, and
is the logical choice for a lectotype.
Solanum troyanum Urb., Symb. Antill. 5: 487. 1908
Type: Type: Jamaica. Trelawney Parish: Mount Diablo, 29 Aug 1905, W. Harris 9000 (lectotype, BM
[BM000849163], designated by Knapp, 2008b: 419;
isolectotypes, K [K000005227], NY [NY00111387]).
Distribution. Endemic to Jamaica, in the Blue
Mountains in forests and forest edges at middle to
high elevations.
Representative specimen. Jamaica, Trelawny Parish,
Proctor 24845 (A, BM, NY, US).
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82
S. Knapp
a
b
c
d
Fig. 6. a, lectotype of Solanum miragoanae Urb., Ekman 7289 (S04-2952) (= S. tetramerum). Reproduced with permission of the Swedish
Museum of Natural History; b, lectotype of S. tetramerum Dunal var. fuertesii O.E. Schulz, Fuertes 284 (NY00111372) (= S. tetramerum
Dunal). Reproduced with permission of the New York Botanical Garden; c, lectotype of S. tetramerum var. luxurians O.E. Schulz, Ekman
H.9979 (S04-2827) (= S. tetramerum Dunal). Reproduced with permission of the Swedish Museum of Natural History; d, lectotype of
S. lobulatum O.E. Schulz, Smith & Smith 1575 (K000196493) (= S. urens Dunal). Reproduced with permission of the Royal Botanic
Gardens, Kew.
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West Indian Solanum
83
Solanum urens Dunal in DC., Prodr. 13(1): 311. 1852
Type: St. Vincent and the Grenadines. St. Vincent:
Bonhomme, A. Anderson s.n. (holotype, G-DC [Morton neg. 8649]).
synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP6 “Structuring the European Research Area” Programme. Reviewer’s comments much
improved the manuscript.
Solanum lobulatum O.E. Schulz in Urb., Symb. Ant.
6: 247. 1909
Type: St. Vincent and the Grenadines. St. Vincent:
Morne Garou, 960 m., H.H. Smith & G.W. Smith
1575 (lectotype, K [K000196493], designated here).
References
Distribution. Only known from montane areas on
the island of St. Vincent, at approximately 1000 m elevation.
Representative specimen. St. Vincent and the Grenadines, St. Vincent, Howard & al. 17711 (BM, NY).
Schulz did not cite a herbarium when describing
Solanum lobulatum, and no sheet of Smith & Smith
1575 is extant at B. The K duplicate of this collection is here designated the lectotype (Fig. 6d) as no
other duplicates have been found, but are likely to
exist. Solanum lobulatum was described from sterile
material, Schulz (1909) stated that the specimen he
examined was annotated “we have only seen one
root”; from the morphology of the specimen at K it is
clear that this is a juvenile individual with a single
stem.
Solanum woodburyi Howard, J. Arnold Arb. 47: 138.
1966
Type: Puerto Rico: S side Luquillo Mountains, El
Toro trail, 2 km from route 191, 13 Oct 1964, R.A.
Howard & G. Taylor 15692 (holotype, A [A00077564];
isotypes, BM [BM000815969], LE, MO [MO638820], NY [NY00111390], US [US-3079749, barcode US00027856]).
Distribution. Endemic to the Luquillo Mountains
in eastern Puerto Rico; known only from the East
Peak region in the Bosque Nacional del Caribe from
150 to 200 m elevation in deep, wooded ravines.
Representative specimen. Puerto Rico, Mun. Luquillo, D’Arcy 1716 (MO).
Acknowledgements
I thank the curators of the herbaria cited in the text for use of
material in their care and for the kind provision of and permission
to use images; Michael Nee (NY) for discussions on locating Solanum type specimens, Pedro Acevedo-Rodriguez (US) for encouragement to work in the West Indies, and Charlie Jarvis (BM)
for discussions on all aspects of typification; this work has been
funded by the National Science Foundation Planetary Biodiversity Inventory Program (award DEB-0316614 ‘PBI Solanum – a
worldwide treatment’ – http:nhm.ac.uk/solanaceaesource) and
my visit to MA from the SYNTHESYS Project http://www.
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Associate Editor: S. Castroviejo
Received: 25-XI-2008
Accepted: 17-II-2009
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